taxonID	type	description	language	source
301187BC2C0DFFF288B0FDE34DDBF835.taxon	materials_examined	Holotype. MNHML MB 29 - 000297, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. Eight spms. MNHML MB 29 - 000298 – 301, stained whole mounts: 3 adults, 1 juvenile. ZMH OL 14518, fixed in Bouin's fluid, preserved in 70 % ethanol: 1 adult, 1 juvenile. ZMH OL 14519, fixed in 70 % ethanol, preserved in 100 % ethanol: 2 spms. Other material. 7 spms, investigated in vivo, preserved in collective sample vials, in the authors' collection.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0DFFF288B0FDE34DDBF835.taxon	etymology	Etymology. Named in honour of Paulo Sousa, soil zoologist, ecologist and ecotoxicologist at the University of Coimbra, research director of the TME experiments.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0DFFF288B0FDE34DDBF835.taxon	diagnosis	Diagnosis. Length <10 mm, less than 40 segments, max. 4 chaetae per bundle, often 3 chaetae per bundle, of same size in a bundle, clitellum girdle-shaped, present also between bursal slits, coelomo-mucocytes with refractile vesicles, nephridia absent at 10 / 11, chylus cells preclitellar with cell canals widened to lacunae, no seminal vesicle, sperm heads 20 – 25 μm, sperm funnel small, spermathecae separate entally, small ectal gland, two sessile diverticula without ciliated subchamber.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0DFFF288B0FDE34DDBF835.taxon	description	Description. Small Fridericia worms. Length 6 – 8 mm (viv), 4.5 – 6 mm (fix); diameter 0.15 mm (viv), 0.16 mm (fix), 0.2 mm at clitellum. Segment number 30 – 35. Chaetae max. 4 per bundle. Formula 2,3,4 – 4,3,2: 2,3,4 – 4,3,2. In preclitellar bundles mostly 4 chaetae, in postclitellar bundles 4 and 3 chaetae, more 3 than 4 in many specimens; two chaetae only in II and in the hindmost 1 or 2 segments. Anteriorly inner chaetae almost as large as outer, posteriorly all chaetae in a bundle of same size. Posterior chaetae slightly larger than anterior (52 – 55: 3 – 3.5 μm and c. 30 – 40: 2 – 3 μm, respectively). Chaetae increasing in size from II to VII (from c. 30 μm to c. 45 μm); chaetae in segments following clitellum as large as preclitellar; caudal chaetae largest (up to 60: 4 μm). Epidermal gland cells one row per segment, at chaetal level, few cells, one each dorsally of lateral chaetae. Body wall c. 15 μm thick, longitudinal muscle layer comparatively thin, 1 – 1.5 x as thick as layer of ring muscles plus epidermis, cuticle very thin, barely visible at 400 x magnification, estimated thickness <0.5 μm. Body surface slightly wavy in about 10 transverse rows per segment. All septa thin. Brain rounded posteriorly, 80: 50 μm, sides almost parallel, anteriorly slightly convex, almost truncate. Pharyngeal glands dorsally connected in IV, V, separate in VI. Dorsal lobes all of same size, ventral lobes of same size in V and VI, small in IV. Oesophageal appendages short, unbranched. Chylus cells in IX – X, canals widened into elongate lacunae. Intestine widened in chylus region. Dorsal blood vessel from XIII. Midgut pars tumida from XX – XXIV, occupying 3 – 3.5 segments lengths. Preclitellar nephridia 4 pairs, from 6 / 7 to 9 / 10, length ratio anteseptale: postseptale about 1: 2; adseptal to medial rise of efferent duct, no terminal vesicle; postclitellar nephridia sparse, often unpaired, first from c. 17 / 18, shape as preclitellar, terminal rise of efferent duct. Coelomomucocytes with small refractile vesicles at periphery, cell length 15 – 20 μm, matrix almost hyaline (viv); lenticytes numerous but inconspicuous (viv), length 5 – 6 μm, i. e. about 1 / 3 of mucocyte length. Clitellum girdle-shaped, cells in c. 25 dense rows, flat (i. e. cells wider than high), conspicuous in vivo, inconspicuous in whole mounts, also present between bursal slits; hyalocytes and granulocytes of same size, diameter 8 – 10 μm (fix). Seminal vesicle absent, developing sperm in anterior half of XI. Spermatozoa c. 80 μm long, heads 20 – 25 μm long. Sperm funnel small, length <1 / 2 body diameter, little longer than wide (e. g. 60: 52 μm, fix), with defined outline, not deformed, collar narrower than funnel body (32 μm). Vas deferens 4 – 5 μm wide (viv, fix). Male copulatory organ: bursal slit longitudinal; male gland small, only slightly projecting into body cavity, not flattened, c. 40 μm long, 30 μm wide, 30 μm high (fix). Subneural glands and other accessory glands absent. Spermatheca: small ectal gland, ectal duct shorter than body diameter, c. 1.5 x as long as ampulla, c. 90 μm long, diameter c. 12 μm, proximal endpiece slightly projecting into ampulla, not widened; two broadly sessile diverticula or diverticula-like protrusions, diameter c. 18 μm, sperm-containing; ampulla tapering proximad, diverticula and ampulla apparently solid (viv) but lumina visible in whole mounts. One mature oocyte at a time, extending over 1.5 – 2 segment lengths.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0DFFF288B0FDE34DDBF835.taxon	discussion	Remarks. Using the tabular comparison of Fridericia species with two spermathecal diverticula in Dózsa- Farkas (2009), F. sousai sp. nov. belongs to a group of 11 species, characterized by separate spermathecae and globular, hemispherical, or short-stalked diverticula. In this group, only F. isseli Rota, 1994 shares the following combination of characters with F. sousai: length <10 mm, less than 40 segments, max. 4 chaetae per bundle, oesophageal appendages short, unbranched, four pairs of preclitellar nephridia, chylus cells preclitellar, coelomocytes with refractile vesicles, seminal vesicle and subneural glands absent, spermatheca with ectal gland. F. isseli differs from the new species in the following traits, among others: (1) chylus cell canals branched, not widened (vs. unbranched, sac-like), (2) clitellum saddle-shaped (vs. girdle-shaped), (3) male gland tripartite with two small accessory glands (vs. accessory glands absent). In the key to Fridericia species in Schmelz (2003), F. sousai would key out together with F. isseli and with F. w a l d e n s t ro e m i Rota & Healy, 1999. The latter differs from F. s o u s a i in having 40 – 54 segments, five preclitellar pairs of nephridia, a small subneural gland in XIV, and proximally fused spermathecae with long-stalked diverticula, among other characters. Figure 5 and Table 2 give a comparison of F. sousai with the other three new species of Fridericia from the same site.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0BFFF188B0FF314C1BFCF1.taxon	materials_examined	Holotype. MNHML MB 29 - 0 0 0 302, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. 60 spms. MNHML MB 29 - 000303 - 311, 9 adult spms, stained whole mounts. ZMH OL 14520, stained whole mounts: 10 spms, 9 adults, 1 subadults. ZMH OL 14521, fixed in Bouin's fluid, preserved in 70 % ethanol: 8 spms. ZMH OL 14522, fixed in 70 % ethanol, preserved in 100 % ethanol: 10 spms. Other material. 134 spms investigated in vivo, preserved in collective sample vials, in the authors' collection.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0BFFF188B0FF314C1BFCF1.taxon	etymology	Etymology. Named in honour of Jörg Römbke, enchytraeidologist, soil ecologist and ecotoxicologist at ECT Oekotoxikologie GmbH Flörsheim, research director of the TME experiments, in grateful recognition of his varied initiatives to support taxonomic work with enchytraeids.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0BFFF188B0FF314C1BFCF1.taxon	diagnosis	Diagnosis. Less than 40 segments, max. 4 chaetae per bundle, clitellum girdle-shaped, cells absent between bursal slits, coelomo-mucocytes without refractile vesicles, nephridia present at 10 / 11, chylus cells post-clitellar, seminal vesicle absent, sperm funnel small, spermathecae joint entally, without ectal gland, ectal duct longer than body diameter, two stalked diverticula with ciliated subchamber and ciliar movement.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0BFFF188B0FF314C1BFCF1.taxon	description	Description. Small Fridericia species. Length 8 – 10 mm when relaxed, 4 – 5 mm when contracted (viv), 6 – 7.5 mm (fix), diameter 0.2 – 0.25 mm (viv), 0.2 – 0.26 at V, 0.28 – 0.36 mm at XII – XIII, 0.22 – 0.29 mm in postclitellar segments (fix). Segment number (29) – 34 – 38 (N = 40), mostly (75 %) 36 – 38. Chaetae max. 4 per bundle, formula 2,3,4 – 4,3,2: (2,3,) 4 – 4,3,2. Posterior 12 – 20 segments with only 2 chaetae per bundle; often all lateral postclitellar bundles with 2 chaetae. Chaetae in caudal segments largest, 65 – 70: 5.5 μm, size of largest anterior chaetae c. 52 – 55: 5.5 μm. Inner chaetae in bundles of 4 always smaller than outer, c. 2 / 3 the size of outer chaetae. Epidermal gland cells (Fig. 2 C, D) in 1 – 3 rows per segment, mid-row at chaetal level, cells pale or yellowish, in regular, alternating pattern, pale cells with clear outline, quasi-rectangular, yellow cells with indistinct outline, smaller; gland cells indistinct or seemingly absent in some specimens. Body wall comparatively thin, 10 – 15 μm thick, longitudinal muscle layer not or only slightly thicker than layer of ring muscles plus epidermis; cuticle very thin (<1 μm), barely visible at 400 x magnification. Septa thin throughout. Brain 100 – 110: 60 – 63 μm, posteriorly truncate, anteriorly slightly convex, sides slightly merging anteriad, almost parallel. Pharyngeal glands connected dorsally in IV, connected or separate in V, separate in VI. Dorsal lobes more or less of same size, ventral lobes strongly increasing in size from IV over V to VI. Oesophageal appendages with few short terminal branches. Chylus cells in 1 / 2 XII – XV, 2 – 2.5 segment lengths; cells in conspicuous longitudinal rows, canals not widened into lacunae. Dorsal blood vessel from XVI – XVII. Midgut pars tumida not distinguished, possibly circumferal. In two consecutive posterior segments, intestinal epithelium with a network of fine canals. Preclitellar nephridia 5 pairs, 6 / 7 – 10 / 11, length c. 125 μm (fix), length ratio anteseptale: postseptale 2: 5, no constriction at septum, medial rise of efferent duct; postclitellar nephridia from 13 / 14. Coelomo-mucocytes pale with blurred vesicles, lenticytes minute, very numerous, optically dominating. Clitellum girdle-shaped, absent between bursal slits, prominent, distinctly higher (c. 5 – 7 x) than non-clitellar epidermis, cells in dense to indefinite rows, hyalocytes and granulocytes alternating, laterally of bursal slits only granulocytes; average diameter of hyalocytes 15 μm, of granulocytes 10 μm, cell height 15 – 20 – (25) μm (fix). Seminal vesicle absent, developing sperm anteriorly in XI. Spermatozoa not numerous on sperm funnel, sperm heads not measurable in vivo, at least 60 μm (fix), probably longer. Sperm funnel small, c. 2 x as long as wide (e. g. 100: 50 μm, or 120: 60 μm, fix), tapering distad, collar about half as wide as funnel body. Vas deferens in wide loops parallel to ventro-lateral body wall, therefore difficult to see, 8 μm wide. Male copulatory organ: bursa longitudinal, male gland c. 80 μm long, 50 μm wide, 40 μm high (fix), kidney-shaped, with thick-walled bursa in concavity. Subneural glands and other accessory glands absent. Spermatheca: no ectal gland; ectal duct c. 250 μm long, longer than body diameter, c. 4 x as long as ampulla, coiled in contracted specimens, diameter 12 μm, with distal swelling up to 20 μm; proximal endpiece (ental bulb) not widened; ampulla with two stalked diverticula oriented ectad, diverticula with ciliated subchamber and rotating spermatozoa in peripheral chamber; ampullae joint entally and with one common attachment to oesophagus dorsally or dorso-laterally. Aspect of joint ampullae plus diverticula varied: common lumen inflated, or collapsed, inner surface smooth or wavy, organs contorted with left ampulla to the right and vice versa, or ampullae compressed in dorso-ventral position with diverticula approached and in staggered position (Fig. 2 A, B). One mature oocyte at a time, occupying up to 3 segment lengths.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C0BFFF188B0FF314C1BFCF1.taxon	discussion	Remarks. Fridericia roembkei sp. nov. belongs to a group of Fridericia species characterized by two spermathecal diverticula with a ciliated subchamber: F. perrieri (Vejdovský, 1878), F. o m e r i Stephenson, 1932, F. rendsinata Dózsa-Farkas, 1972, F. u l r i k a e Rota & Healy, 1999, F. h e a l y a e Schmelz, 2003, F. dozsae Schmelz, 2003 (Schmelz 2003: 296, 341 f.), and partly also F. galba (Hoffmeister, 1843). It differs, together with F. m a rg i na ta described below, from all species of this group in the proximal fusion of the spermathecal ampullae. Using the tabular comparison of Fridericia species with two spermathecal diverticula in Dózsa-Farkas (2009), F. roembkei sp. nov. belongs to a group of 10 species with proximally fused spermathecae. It differs from all species of this group in the absence of spermathecal ectal glands, in an exceptionally long spermathecal ectal duct, and in the ciliation of the diverticula. Differences to F. marginata sp. nov., a remarkably similar species, are dealt with in the remarks section of that species, see also Figure 5 and Table 2. The aspect of the spermathecae in F. roembkei varies considerably. The ampullae are often compressed, twisted, and in upright position, as shown in Fig. 2 A, or they are inflated into a spherical chamber — much more than shown in Fig. 2 B — mimicking the aspect in F. gamotheca (Fig. 2 F). Both aspects can be found at different times in the same living specimen. These unusual variations may be related to the thin and apparently soft ampullar walls. In the key to species of Fridericia in Schmelz (2003), F. roembkei would key out together with F. g a m o t h e c a Issel, 1905. F. gamotheca as originally described (Issel 1905) and redescribed (Rota 1995) from several Italian localities is distinguishable from F. roembkei by the more oval spermathecal diverticula, a shorter spermathecal ectal duct (1.5 x times as long as the ampulla), a larger sperm funnel, and higher segment number (40 – 44). Furthermore, diverticula are not ciliated (Schmelz pers. obs.) and the inflated aspect of the joint ampullae is a constant feature, not a transitory state as in F. roembkei. Some further possible differences are not straightforward because of variations noted in Rota (1995) regarding shape of diverticula, seminal vesicles and spermathecal ectal glands. Another variant of F. gamotheca from Morocco of uncertain taxonomic status (Schmelz 2003: 193 f.) has spherical diverticula, only 28 – 33 segments (Dózsa-Farkas 1989), and a ventral clitellum as described for F. roembkei: girdle-shaped, absent between bursal slits (Schmelz 2003). It differs from F. roembkei in a short and thick ectal duct and in the presence of spermathecal ectal glands. The morphological variability documented in the redescriptions of Dózsa-Farkas (1989) and Rota (1995) have led Collado et al. (2012) to assume that F. g a m o t h e c a is a species complex. Remarkable in F. roembkei is further the presence of both pale and yellow epidermal gland cells (Fig. 2 C, D), distinct only in living specimens, and clearly representing two different cell types. The pale cells are quasirectangular, filled with pale vesicles and mostly arranged in one row at chaetal level, while the yellow cells are smaller spots of irregular outline, not arranged in one row but nonetheless distributed with some regularity.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C09FFFF88B0FC2A485DFA5A.taxon	materials_examined	Holotype. MNHML MB 29 - 000312, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. 15 spms. MNHML MB 29 - 000313 – 315, stained whole mounts, 2 adults, 1 subadult. ZMH OL 14523: 4 juvenile spms, stained whole mounts. ZMH OL 14524, fixed in Bouin's fluid, preserved in 70 % ethanol: 8 spms. ZMH OL 14525, fixed in 70 % ethanol, preserved in 100 % ethanol: 5 spms. Other material. 37 spms investigated in vivo, preserved in collective sample vials, in the authors' collection.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C09FFFF88B0FC2A485DFA5A.taxon	etymology	Etymology. Named for the ventro-lateral margins of the clitellum, consisting only of granulocytes.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C09FFFF88B0FC2A485DFA5A.taxon	diagnosis	Diagnosis. Less than 40 segments, max. 4 chaetae per bundle, clitellum saddle-shaped with ventral margins consisting of granulocytes only, coelomo-mucocytes with refractile vesicles, nephridia present at 10 / 11, chylus cells post-clitellar, no seminal vesicle, sperm heads 90 μm, sperm funnel small, spermathecae joint entally, without ectal gland, two stalked diverticula with ciliated subchamber, ampulla and diverticula elongate.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C09FFFF88B0FC2A485DFA5A.taxon	description	Description. Body colour opaque grey-white, often with a yellowish tinge (viv), slow body movements. Length c. 7 – 8 mm (viv), diameter c. 0.3 mm (viv). Segments 35 – 39. Chaetae max. 4 per bundle, formula 3,4 – 4,3,2: (3), 4 – 4,3,2. Ventrally 2 per bundle from XXV to posterior end. Inner chaetae shorter than outer. Chaetae in caudal segments largest, 65 – 70: 5.5 μm, size of largest anterior chaetae c. 55: 4.5 μm. Epidermal gland cells scarce, pale, one row at chaetal level, cells quadrangular or without clear outline. Body wall c. 20 μm thick with longitudinal muscle layer thickest (c. 15 μm), cuticle thin (<1 μm). Preclitellar septa 4 / 5 – 9 / 10 thickened, thinner at 10 / 11 ff. Brain rounded posteriorly, lateral sides parallel. Pharyngeal glands all united dorsally, dorsal lobes slightly increasing in size from IV to VI, ventral lobes strongly increasing in size from IV to VI. Oesophageal appendages elongate with 2 – 3 short or elongate branches in distal third. Chylus cells in XIII – XIV, 1.5 – 2 segment lengths, canals not widened into lacunae. Dorsal blood vessel from XVI – XVII. Pars tumida of midgut in XXVI – XXIX, extending over 2 – 4 segments, not distinguished in all specimens. Preclitellar nephridia 5 pairs, 6 / 7 – 10 / 11, length ratio anteseptale: postseptale 2: 3, medial to subterminal rise of efferent duct, terminal vesicle absent. Postclitellar nephridia with subterminal to terminal rise of efferent duct and elongate postseptale. Coelomo-mucocytes broadly oval, length 25 – 35 μm, with refractile vesicles at periphery in one incomplete row, vesicle diameter c. 1 μm, matrix transparent; cells very numerous, filling all spaces, making body opaque with grey-yellow tinge (viv); lenticytes minute, length c. 6 – 8 μm, sparse. Clitellum saddle-shaped, prominent, cells in c. 35 dense rows with granulocytes and hyalocytes alternating; mid-ventral aclitellar field as wide as distance between bursal slits; hyalocytes absent at ventro-lateral borders and in a semi-circular field laterally of and around bursal slits; here only granulocytes in indefinite rows or in reticulate arrangement. (Single hyalocytes near ventral border between male and female pores observed in one specimen). Seminal vesicle absent; occasionally dense masses of developing sperm in anterior half of XI, and septum 10 / 11 bulged forward dorsally, unpaired. Spermatozoa> 200 μm long, heads c. 90 μm, not numerous on top of sperm funnel (Fig. 3 C). Sperm funnel comparatively small, hardly visible among coelomocytes, developing sperm, and oogonia (viv), about half as long as body diameter, tapering proximally, c. 2.5 x as long as wide (e. g. 140: 60 μm), collar not wider than funnel body, not folded outwards. Vas deferens in dense irregular coils ventro-laterally, 8 μm wide. Male copulatory organ elevating body suface even when completely retracted (viv), transverse copulatory muscles strongly developed. Male glands broadly oval in ventral view, flat in side view, 1.5 x as long as wide (e. g. 120 μm long, 80 μm wide, 50 μm high) bursal slits longitudinal, bursa thick-walled, with small and flat modiolus (comp. Schmelz 2003: 49) around ectal opening of vas deferens (the primary male pore); bursa plus male gland almost spherical. Subneural glands and other accessory glands absent. Spermathecae: no ectal gland; ectal duct about as long as body diameter, slightly widened in distal part, tapering again near ectal pore, ental bulb not projecting into ampullar lumen, only slightly wider than ectal duct, ectal duct canal straight throughout. Ampullae 1.5 – 2 x as long as wide, tapering entad, ental tips fused, creating a common V-shaped lumen. Ampullar walls 4 – 6 μm thick, uniform, no histological differences observed, with smooth outer and inner surface, inner surface occasionally wavy, as a transitory state during contraction. Diverticula inserting on opposite sides of ental bulb at base of ampulla and projecting obliquely ectad, distance (= maximum width of spermatheca) c. 80 μm, c. 1 / 3 body diameter. Each diverticulum 2 x as long as wide, diameter c. 20 μm, lumen subdivided into peripheral spermcontaining chamber and ciliated sub-chamber; both chambers connected by a porus; peripheral chamber wider than long, with dense sperm roll circulating; sub-chamber longer than wide, with ciliar movement, widely connected with ampullar lumen. No ciliar movement in ampulla. One mature oocyte at a time, extending over c. 2 segment lengths.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C09FFFF88B0FC2A485DFA5A.taxon	discussion	Remarks. The new species owes its name to the saddle-shaped clitellum with ventro-lateral margins consisting only of granulocytes, a character new in Fridericia, but common in some other genera, e. g. Guaranidrilus, Hemienchytraeus, Achaeta, Marionina. In the other Fridericia species with saddle-shaped clitellum the ventro-lateral borders do to our knowledge not differ from the rest and are composed of granulocytes and hyalocytes. However, not all species of the genus have been examined for that character. A further peculiarity is the slenderness of ampulla and diverticula, both about twice as long as wide. Using the tabular comparison of Fridericia species with two spermathecal diverticula in Dózsa-Farkas (2009), F. marginata sp. nov. belongs to a group of 10 species characterized by proximally fused spermathecae. None of these species has the character combination of max. 4 chaetae per bundle, saddle-shaped clitellum, and stalked spermathecal diverticula. Using the key to Friderica species in Schmelz (2003), F. marginata would key out together with F. gamotheca Issel, 1905. This species has also coelomocytes with few refractile vesicles, as F. marginata (Rota 1995, Schmelz pers. obs.). It differs from F. marginata in larger body size and higher segment number (37 – 50 according to Rota 1995), in a girdle-shaped clitellum according to Schmelz and Collado (2010), and in a complete fusion of the two spermathecal ampullae. A variant of F. gamotheca described from Morocco (Dózsa-Farkas 1989) and of uncertain taxonomic status (Schmelz 2003) is more in the size range of F. m a rg i na ta (length 5 – 7 mm, 28 – 33 segments), but differs in a girdle-shaped clitellum, coelomocytes without refractile vesicles and in the presence of a spermathecal ectal gland, among other characters. F. gamotheca is a problematic species, see F. roembkei sp. nov., remarks. F. m a rg i n a t a and F. roembkei spp. nov. are quite similar at first sight, but they are easily distinguished by the coelomocytes — with refractile vesicles in F. m a rg i n a t a and without them in F. roembkei — and by the ventral pattern of the clitellum, see Figure 5. Further differences are more difficult to observe and may not be evident unless specimens of both taxa are available for comparison (in the following, character states of F. marginata first): spermathecal diverticula and ampulla elongate / not elongate; spermathecal ectal duct not longer / longer than body diameter; yellow epidermal gland cells absent / often present; pharyngeal glands in VI united / separate dorsally (Table 2).	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	materials_examined	Holotype. MNHML MB 29 - 000316, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. 13 spms. MNHML MB 29 - 000317, – 318, stained whole mounts, 1 adult, 1 subadult. ZMH OL 14526, 5 adults, unstained whole mounts. ZMH OL 14527, fixed in Bouin's fluid, preserved in 70 % ethanol: 9 spms. ZMH OL 14528, fixed in 70 % ethanol, preserved in 100 % ethanol: 2 spms. Other material. 11 spms investigated in vivo, preserved in collective sample vials in the authors' collection.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	etymology	Etymology. Named for the ciliar movement in the spermathecal ampulla.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	diagnosis	Diagnosis. More than 50 segments, max. 4 chaetae per bundle, clitellum saddle-shaped, coelomo-mucocytes without refractile vesicles, nephridia present at 10 / 11, chylus cells post-clitellar, no seminal vesicle, sperm heads c. 120 μm, sperm funnel small, spermathecae separate entally, no ectal gland, two stalked diverticula with ciliated subchamber, ampulla constricted ectally, with ciliar movement near ental bulb.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	description	Description. Worms with transparent and opaque parts and often greenish gut (viv), slow body movements. Length 13 – 15 mm (viv), 10 – 12 mm (fix), diameter 0.30 – 0.35 mm at XII – XIII, 0.25 – 0.28 mm in rest of body (viv, fix). Segment number 55 – 59 in fully adult worms (N = 10), often less in subadults. Chaetae max. 4 per bundle; formula 4 – 4,3,2: 4 – 4,3,2. Mostly 4 chaetae per bundle in anterior body half (e. g. down to XXVII in a specimen with 58 segments) and 2 chaetae per bundle in posterior body half, 3 chaetae only in bundles of a few mid-body segments. Anterior chaetae max. 65 μm long and c. 5 μm wide, inner chaetae c. 3 / 5 the size of the outer, laterals slightly smaller (e. g. 56 μm) than ventrals; postclitellar chaetae not reduced in size, chaetae in caudal segments not longer than outer chaetae in anterior segments, but slightly thicker (up to 6 μm). Single packages of detached chaetae occasionally present in body cavity. Epidermal gland cells pale, quasi-rectangular, in 3 rows in anterior segments, most cells in mid-row at chaetal level, mid-dorsally and mid-ventrally no cells; in some specimens cells inconspicuous or apparently absent. Body wall 20 – 30 μm thick, cuticle thin (<1 μm), preclitellar septa not conspicuously thickened. Brain egg-shaped (viv), c. 100: 70 μm (fix), rounded on all sides, anteriorly with very slight convexity, almost truncate. Pharyngeal glands dorsally united in IV, V, separate in VI; dorsal and ventral lobes increasing in size from IV to VI; in IV ventral lobes separate from dorsal lobes. Oesophageal appendages elongate with few terminal branches, extending to VI. Chloragocytes with greenish or greyish vesicles. Chylus cells in XV – XIX, occupying 3.5 – 4 segment lengths, canals not widened into lacunae. Dorsal blood vessel from XXIII – XXIV. Midgut pars tumida not seen. In two consecutive posterior segments, intestinal epithelium with a network of fine canals. Preclitellar nephridia 5 pairs, from 6 / 7 to 10 / 11, constricted at septum, length ratio anteseptale: postseptale between 3: 4 and 2: 3, adseptal to medial rise of efferent duct, no terminal vesicle. Postclitellar nephridia from 13 / 14, constriction less marked, also with large anteseptale and subterminal rise of efferent duct, no terminal vesicle. Coelomo-mucocytes small, length 20 – 30 μm (fix), pale, 1.5 – 2 x as long as wide, matrix with strongly blurred vesicles, almost hyaline, nucleus distinct, with 1 or 2 minute nucleoli; margins of cells in oblique position appear dark (light refraction); coelomo-lenticytes 7 – 8 μm long. Both cell types numerous but not filling body cavity. Cell aggregations with grey tint, cells appear to stick to one another. Clitellum saddle-shaped with cells in c. 35 dense, transverse rows, conspicuous but not much elevated, cell height c. 15 μm, 20 μm in contracted worms (fix), granulocytes and hyalocytes alternating, diameters 8 and 15 μm, respectively (fix); hyalocytes extending down to ventro-lateral margins; female pores surrounded by granulocytes only. Mid-ventral aclitellar field wide at level of bursal slits, extending laterally beyond slits; in the rest, aclitellar field narrower than distance between ventral chaetal bundles, narrowest at level of female pores. Seminal vesicle absent, developing sperm in anterior half of XI. Spermatozoa> 300 μm long, heads c. 120 μm. Sperm funnel conical / cylindrical, variable in shape and often of irregular outline, about half as long as body diameter, collar narrower than funnel body, with few spermatozoa attached. Vas deferens 20 μm wide (fix) proximally near sperm funnel, continuously tapering to 9 μm distally (fix) near male gland. Male copulatory organ: male gland distinctly longer than wide, e. g. 120 μm long, 70 μm wide, 50 μm high (fix), bursal slit H-shaped. Subneural glands and other accessory glands absent. Spermatheca: no ectal gland; ectal duct about as long as body diameter, c. 18 μm wide (viv), not widened in distal part; ental bulb not projecting into ampullar lumen, only slightly wider than ectal duct, ectal duct canal straight throughout; ampullae with widely separate attachment to oesophagus dorso-laterally; each ampulla 1.2 – 1.5 x as long as wide, rounded entally and constricted ectally at transition to diverticula; walls 4 – 6 μm thick, uniform, no histological differences observed, with smooth outer and inner surface, inner surface somewhat uneven, occasionally wavy, as a transitory state during contraction. Diverticula inserting on opposite sides of ental bulb at base of ampulla and projecting obliquely ectad, distance (= maximum width of spermatheca) c. 60 – 70 μm, c. 1 / 4 body diameter. Each diverticulum c. 1.4 x as long as wide, diameter c. 20 μm, lumen subdivided into peripheral sperm-containing chamber and ciliated sub-chamber; both chambers connected by a porus; peripheral chamber wider than long, with dense sperm roll circulating; sub-chamber longer than wide, widely connected with ampullar lumen. Ciliar movement in peripheral chamber of diverticula and also in distal part of ampulla, in vicinity of ental bulb surface (Fig. 4 B, arrow).	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	discussion	Remarks. F. ciliotheca belongs to the group of Fridericia species with spermathecal diverticula that have a ciliated subchamber. The movements of the cilia keep the packages of allosperm in constant rotation, probably a means to maintain them in viable state. Additionally, ciliar movement was also observed inside the ampulla, in immediate vicinity of the ental bulb (Fig. 4 B, arrow). The species owes its name to this peculiarity, observed here for the first time in Fridericia (and perhaps Oligochaeta in general, since to our knowledge Fridericia is the only oligochaete taxon where ciliated spermathecae occur). The character is only seen in living specimens. Insertion and length of the cilia could not be ascertained, but a continuity with the cilia in the diverticular subchamber was not evident, either. It seems, therefore, that the cilia insert in the ampullar epithelium. Among Fridericia species with ciliated subchamber, most similar to F. ciliotheca sp. nov. is F. u l r i k a e Rota & Healy, 1999, with a maximum of 4 chaetae per bundle, saddle-shaped clitellum, long spermatozoa, elongate male glands, and separate spermathecae. Apart from the absence of cilia in the ampulla, F. u l r i k a e differs from F.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	description	ciliotheca in the following characters: 4 pairs of preclitellar nephridia (vs. 5), seminal vesicle always large, extending over two segment lengths (vs. small, confined to XI), pharyngeal glands in VI connected dorsally (vs. not connected), spermathecae distinctly larger, with diverticula 40 – 50 μm wide (Rota & Healy 1999, Schmelz pers. obs.) (vs. 20 μm), ectal duct distinctly shorter than body diameter (vs. as long as body diameter). Figure 5 and Table 2 give a comparison of F. ciliotheca with the other three new species of Fridericia from the same site. Achaeta coimbrensis sp. nov. (Fig. 6 A – E)	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	materials_examined	Holotype. MNHML MB 29 - 0 0 0 319, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), crop site (Table 2); IV 2010. Paratypes. MNHML MB 29 - 000320 – 321, two spms, adult, subadult, stained whole mounts, same data as for holotype. Other material. No further material available.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	etymology	Etymology. Named after the city of Coimbra, Portugal.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	diagnosis	Diagnosis. Length c. 2 mm, 18 – 21 segments, no pyriform or lentiform glands, oesophageal appendages in V only, no secondary pharyngeal gland lobes, dorsal blood vessel from 1 / 2 VII, abrupt widening of intestine at 1 / 2 VII, two pairs preclitellar nephridia at 6 / 7, 7 / 8, gonadal region shifted one segment forward, clitellum with narrow dorsal and wide ventral interruption, hyalocytes on dorsal body half only, in irregular, not baguette-shaped longitudinal rows, no seminal vesicle, sperm funnel and male copulatory organs small, male gland tripartite, with two small accessory glands lining larger central gland, spermathecae short, ectal pores lateral.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	description	Description. Body length c. 2 mm (holotype 2.16 mm), diameter c. 0.1 mm (holotype: 0.085 mm at V, 0.11 mm at XI – XX) (fix). Segment number 18 (holotype), 20 (subadult paratype), 21 (adult paratype). Head pore on prostomium, a longitudinal slit. Body wall very thin, thickness 2 – 3 μm, layers (cuticle, epidermis, ring muscles, longitudinal muscles) barely distinguishable. No pyriform glands. No lentiform glands seen. Prostomium wider than long, with thickened frontal tissue protruding into lumen, reducing free coelomic space inside prostomium, no frontal recess observed (viv, fix); at least two pairs of papillae distinguishable. Brain broadly pear-shaped, length 50 μm, width 30 μm, with a minute pair of prostomial ganglia. Ventral nerve cord ganglionate from V on, II – IV fused into suboesophageal ganglion. Ganglia at mid-length with large nonstaining cells ventrally (Fig. 6 B). Two pairs of post-pharyngeal ganglia; the outer pair elongate, on afferent fascicles of pharyngeal glands. Septa 4 / 5 – 6 / 7 slightly thickened. Pharyngeal glands in IV – VI, all with ventral lobes, dorsal lobes united in IV and V, in VI united or separate, ventral lobes may be separate from dorsal lobes here. Secondary pharyngeal gland lobes absent. Oesophageal appendages in anterior half of V, a pair of spherical dorso-lateral bodies, lobe diameter almost equals that of dorsal pharyngeal gland lobes. Preclitellar nephridia at 6 / 7 and 7 / 8; anteseptale as long as wide, postseptale c. 2 – 3 x as long as anteseptale, gradually narrowed into short efferent duct; no terminal vesicle. Postclitellar nephridia not more than 1 – 3 altogether, at 15 / 16 or 16 / 17 only, paired or unpaired, shape as preclitellar. Chloragocytes in one specimen with large yellow-brown vesicles, here over the entire body; coverage of intestine incomplete; first cells in IV; many chloragocytes floating freely in the coelom. Dorsal blood vessel from 1 / 2 VII. Gut abruptly widening in 1 / 2 VII, immediately behind dorsal blood vessel origin, with small inner recess of intestinal epithelium directed anteriad. Pars tumida of midgut in XII – XIV or XIII – XV, cell height more than half the gut diameter (Fig. 6 B). Two types of coelomocytes: type I c. 25 µm long, flat, pale, inconspicuous, slightly longer than wide, with pointed projections at narrow ends; type II c. 15 µm long, filled with distinct pale vesicles. As a third type of cells in the coelom, detached chloragocytes. Coelomocytes pale and inconspicuous, chloragocytes very conspicuous (viv). Testis and sperm funnels in X, male efferent apparatus and ovary in XI. Clitellum in XI – 1 / 2 XII, not developed mid-dorsally and mid-ventrally, mid-dorsal gap narrow, mid-ventral gap as wide as distance between male pores. Glands in 16 – 18 transverse rows; hyalocytes in four longitudinal rows, one dorsal and one dorso-lateral on each side; dorsolateral rows somewhat irregular in posterior half (Fig. 6 C); rows not compacted, hyalocytes not protruding into coelom beneath body muscles. Testis unpaired mid-ventrally. Sperm developing freely in X, no seminal vesicle. Sperm heads c. 14 µm long (viv), total sperm length not measured. Sperm funnel comparatively small, about 1 / 4 as long as body diameter or less, about 2 x as long as wide, barrel-shaped, oval in diameter, so length-width ratio larger in side view (1.5: 1); collar narrower than funnel body. Vas deferens 3 – 4 μm wide, thinnest in mid-section, widening ectally near male pore, irregularly coiled entally near sperm funnel. Male pores in XI, widely paired, on body surface, at mid-length of clitellum; male glands multiple, arranged longitudinally: one larger compact horse-shoe shaped gland surrounding male pore, 30: 25 μm, and 1 – 2 small accessory glands attached anteriorly and posteriorly, diameter <10 μm. Spermathecae with lateral ectal pores, blind-ending, confined to V, shorter than body diameter, projecting entad as a stiff finger or attached to inner body wall, length c. 60 μm, diameter c. 8 – 10 μm, ental end club-shaped, diameter 15 μm, sperm at about 2 / 5 length ectally, ectal duct canal lined with cuticle for only 8 μm. One mature oocyte at a time, filling out one segment.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C07FFF988B0FA49480FFD61.taxon	discussion	Remarks. The new species belongs to a group of Achaeta species without pyriform glands and with lateral spermathecal pores. Within this group, a forward shift of the sexual organs (except the spermathecae) by one segment is characteristic of the European species A. pannonica Graefe, 1989, A. diddeni Graefe, 2007, and the South American A. hanagarthi Schmelz, 2008. Among these three species, A. diddeni is most similar to the new species. Both have in common: small body size, low segment number (22 – 24 in A. diddeni), oesophageal appendages in V only, dorsal blood vessel from VII, two pairs preclitellar nephridia at 6 / 7, 7 / 8, clitellum open dorsally and ventrally, no ' baguette' - shaped concentration of hyalocytes dorso-laterally, male reproductive system and spermathecae small. Differences between both species are as follows: A. diddeni has (1) six segmental epidermal lentiform gland cells (vs. none seen), (2) secondary (post-septal) pharyngeal glands in V and VI (vs. none), (3) the intestine widens gradually (vs. abruptly at 1 / 2 VII), and (4) accessory male glands are absent (vs. present). A. pannonica has the same pattern of similarities and differences, except for the nephridia (3 preclitellar pairs, vs. 2) and lentiform glands (more than 6 per segment, vs. none seen). The South American A. hanagarthi is also without secondary pharyngeal gland lobes but differs in segment number (> 30), preclitellar nephridia (1 pair, at 7 / 8), and spermathecae (long, extending into VI or VII), among other characters. Furthermore, A. coimbrensis sp. nov. appears to be smaller than the three species, but dimensions of the new species are from fixed material. Lentiform epidermal gland cells were not found after intensive observation of living and preserved specimens, but their absence cannot be stated with certainty because these glands are sometimes difficult to see. — The variability of the pharyngeal gland lobe pattern in VI may be artifactual in parts (the adult paratype specimen is damaged in this region) and cannot be assessed here taxonomically. — The non-staining cells inside the ventral nerve cord are common in Achaeta species, but their function is still unclear. They may be neurosecretory gland cells. — The low number of type specimens is unsatisfactory, but we hope to add more material in future surveys at the type locality.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C01FFE588B0FD5A481AFC01.taxon	materials_examined	Holotype and paratypes. Not designated in original description, not located, probably lost. Investigated material. Three spms, 1 adult, 2 early-stage subadults. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Agricultural School (ESAC), crop site (Table 2); IV 2010. One of the subadults cut in halves: posterior half used for DNA extraction and sequence generation; anterior half as voucher in 100 % ethanol. Rest fixed in Bouin's fluid, stained with paracarmine, whole-mounted in Canada balsam.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C01FFE588B0FD5A481AFC01.taxon	description	Description. Dimensions are from the single adult whole-mounted specimen, unless stated otherwise. Specific traits found only in one of the whole-mounted specimens are marked with " AD " (adult specimen) or " SA " (subadult specimen). Differences of the original description are inserted, marked by " NC ". Length c. 5 mm (viv), 2.5 – 4 mm (fix); diameter (fix) 0.22 mm at V, XX, 0.26 mm at XII. Segments 25 (N = 2), NC: (27) – 31 – (35). Chaetae straight with weak ental hook, 2 and 3 per bundle, ventrally only 3. Formula 2,3 – 2,3: 3 – 3. In lateral bundles 2 chaetae mostly near clitellum. Chaetae absent at XII. Lateral chaetae in truly lateral position, not shifted dorsally. Chaetae mostly 40 – 50 µm long and 3 – 4 µm wide, largest in caudal segments (> 50 μm), smallest in II (c. 35: 3 µm); size differences between ventral and lateral chaetae not ascertained. Loose packages of detached chaetae present in the coelom in the adult specimen. Head pore mid-dorsally at 0 / I, a longitudinal slit, length c. 15 µm. Prostomium rounded, broadly conical, about as long as wide or high, shorter in fixed specimens. Epithelium without frontal recess, increasing in height towards the front, lined basally with musculature (not shown in Fig. 7 A). No sensory papillae apparent (fix), no prostomial papillae seen. Fibers of subepithelial muscular lining extending across lumen of prostomium in various longitudinal, transverse and oblique directions (not shown in Fig. 7 A). Peristomium c. 3 / 5 the length of following segments (II, III etc.). Epidermal gland cells not observed. No segmental dorsal pores. Body wall thin, c. 10 µm, up to 20 µm in anterior segments. Longitudinal muscle layer about as thick as layers of ring muscles, epidermis, and cuticle together. Cuticle very thin, ring muscles in irregular arrangement (optical section). Septa very thin, slightly thickened at 4 / 5 – 6 / 7. Brain 85: 60 µm (fix), rounded or truncate posteriorly, sides converging anteriad, anteriorly concave; prostomial nerves with a pair of ganglia (SA), or ganglia not seen (AD). Ventral nerve cord medullar, slightly constricted and with fewer perikarya near the septa. Inner pair of postpharyngeal ganglia situated on top of oesophageal appendages; outer pair on afferent fascicles, enlarged and apparently with pharyngeal gland tissue in adult specimen (Fig. 7 A, arrow). Pharyngeal pad comparatively large, 130 µm long and 90 µm wide in adult specimen (fix), heavily invested with musculature dorsally, fibers extend backwards into V. Oesophageal appendages in III – IV, a pair of short, unbranched, free-floating, sac-like tubes with wide lumen, attached separately and adjacently to oesophagus dorsally behind pharyngeal pad and below post-pharyngeal ganglia, extending obliquely backwards and downwards, blind-ending ventrally below pharyngeal gland lobes of IV (AD); maximum diameter 40 µm, estimated length c. 160 µm; epithelium 3 – 4 µm thick, not folded, smooth towards coelom, slightly rugose towards inner lumen. Pharyngeal glands in IV – VI, decreasing in size from IV to VI (AD) or of equal size in IV – VI (SA); dorsal lobes separate or with narrow dorsal connection, ventral lobes present from IV – VI, no secondary glands. Intestinal diverticula in VII, dorsally and laterally of oesophagus, with common unpaired opening into oesophagus dorsally, lateral wings widely connected dorsally of oesophagus, laterally flattened, extending forwards and backwards to the same degree; diverticular epithelium smooth, not folded, lined with blood sinus and chloragocytes, with histological properties different from adjacent oesophageal and intestinal epithelium: somewhat yellowish in whole mounts, not staining with paracarmine. Intestine widening strongly from c. 50 µm diameter at 6 / 7 to c. 150 µm in VIII; widening abrupt at 7 / 8 in contracted specimens. Pars tumida of midgut not distinguished. Intestine filled with amorphous humous substance mixed with sand grains up to 40 µm in diameter. Chloragocytes with dense layer from VI, covering also intestinal diverticula; cell diameter c. 10 µm (fix); in postclitellar segments cells often higher than wide (fix). Blood colourless, dorsal blood vessel from VII, large and pulsating, bifurcating in I below brain into circumesophageal connectives, these uniting into ventral vessel in IV. Two pairs of commissural vessels seen in III posteriorly, branching off dorsal vessel close to each other. Preclitellar nephridia 6 pairs, from 4 / 5 to 9 / 10. Postclitellar nephridia from 13 / 14, absent at several positions around XVII – XX; length c. 80 µm; anteseptale with funnel only, postseptale laterally flattened, c. 1.5 x as long as high, with well-developed interstitial tissue, efferent duct about half the length of postseptale, rising terminally, nephroporus with small terminal vesicle, conspicuous in vivo. Coelomocytes one type, mucocytes, flat, broadly oval or broadly pear-shaped, c. 20 µm long (viv), 16 – 22 µm long and 10 – 12 µm wide (fix), filled with small, slightly refractile vesicles (viv), nucleus and vesicles distinct (fix). Cells numerous, but not in dense aggregations. Clitellum in XII and XIII (NC: XII – 1 / 2 XIII), extending beyond chatae of XIII by 2 – 3 transverse cell rows; girdle-shaped, cells in c. 32 dense rows, hyalocytes evenly distributed on all sides, present also mid-ventrally; celldiameter c. 8 – 10 µm (fix), cells higher than wide laterally (c. 15 µm, fix). Testes and sperm funnels in XI, male pores and ovaries in XII. Testes and developing sperm enclosed by a common membrane, i. e. sperm developing in testis sacs and not freely in the coelom. Two testis sacs present, one on each side, not lobed, confined to lateral sides of XI, occupying up to one segment length. Spermatozoa c. 50 µm long, heads c. 12 µm (fix); lengths in vivo not measured. Sperm funnel small, 1.5 – 2 x as long as wide (e. g. 60: 36 μm, fix), tapering distad, collar as wide as funnel body. Vas deferens short (estimated length c. 400 µm), not winded into coils, proximal region with wide ciliated lumen, tube diameter here c. 20 μm, tapering distad more or less continuously down to 7 – 8 µm near male pore. Tube walls c. 2 µm thick throughout, cilia in lumen conspicuous. Male copulatory organ with bursa, gland, and copulatory muscles. Gland rounded and compact, c. 50 μm long, 40 μm wide, 30 µm high (fix), bursal slit longitudinal, c. 25 µm long, transverse copulatory muscles only in immediate vicinity of male gland. Subneural glands absent. Spermathecae as simple, mid-dorsally united tubes. Each tube with a double swelling, the ectal one possibly representing ectal duct, but cuticular lining not distinguished here and sperm present in both swellings. Ental swelling wider (c. 25 µm) than ectal swelling (c. 20 µm). No diverticula, no ectal glands. (NC: Ectal duct long and narrow, with distinct canal, sometimes with few separate gland cells near ectal pore). Ectal porus minute, at halflength between lateral chaetae of IV and V, slightly shifted ventrally. Two mature eggs in the adult specimen, each occupying about one segment length (NC: 1 – 8 eggs at a time). No intestinal or coelomic parasites observed.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
301187BC2C01FFE588B0FD5A481AFC01.taxon	discussion	Remarks. Enchylea heteroducta had been described from laboratory cultures of unknown origin (Nielsen & Christensen 1963). This is the first record of the species from a natural habitat; the note in Schmelz and Collado (2012) refers to the same record. Specimens underlying the original description had more segments (27 – 35), up to 8 mature oocytes at a time, and a slightly different spermatheca: There was an ectal duct at the same place where we found a sperm-containing swelling. For the time being we assume that these differences are intraspecifi c. Several structures differ in appearance between the adult and the subadult specimen; more material is necessary to decide which character state is the ' valid' one. The reduced size of the hindmost pair of pharyngeal glands in the adult specimen may be a case of degeneration, although it agrees with the original description. Peculiar in this species are the combined presence, position and shape of oesophageal appendages and intestinal diverticula, and the short and stout vasa deferentia. Gut appendages are among the most important characters to distinguish genera among enchytraeids, therefore the new species was accomodated in a genus of its own. The oesophageal appendages are very similar to those in the Enchytraeus albidus group sensu Schmelz and Collado (2010), the only difference in the latter being their smaller size relative to the rest of the body and a thicker epithelium relative to the lumen diameter. Enchytraeus differs from Enchylea in the absence of intestinal diverticula, among other characters, and species of the E. albidus group are much larger than Enchylea, with body lengths around 3 cm. The intestinal diverticula in E. heteroducta are almost unique in the family: paired in VII laterally, broadly interconnected dorsally and with one common opening into the intestine. Perhaps they are better described in the singular form, as one dorsal diverticulum, bifurcating twice, first in lateral, then in antero-posterior direction. Such a structure is also described in Guaranidrilus sawayai Righi, 1973, the only species in this predominantly neotropical genus where the diverticula are not separate. Guaranidrilus differs from Enchylea in the oesophageal appendages (rounded and compact bodies in VI, when present), in not more than 2 chaetae per bundle and in free spermathecae, among other characters. Crop site Meadow site * Enchytraeus bulbosus Nielsen & Christensen, 1963 x x * Enchytraeus dichaetus Schmelz & Collado, 2010 x x = Enchytraeus christenseni bisetosus Rota & Healy, 1994 1 Form without refractile granules in coelomocytes (Schmelz & Collado 2010), possibly comprising several species. 2 Form with straight canal of spermathecal ectal duct near ampulla (Schmelz 2003: 291 f.). 3 Possibly new species but without reference material of sufficient quality. 4 Form with refractile granules in coelomocytes (Schmelz & Collado 2010), possibly comprising several species. 5 Form with spiral canal of spermathecal ectal duct near ampulla (Schmelz 2003: 291 f.). 6 The recent split of M. argentea into four species (Rota 2012) could not be considered here. Nielsen and Christensen (1963) suggested an affinity of Enchylea to Enchytraeus, because of the shape and position of the oesophageal appendages. The presence of testis sacs, one of the traits not mentioned originally but present in our material, supports their assumption. Testis sacs are known so far only in Enchytraeus and in Lumbricillus, the latter without oesophageal appendages. These two genera further agree with Enchlyea in the short nephridial anteseptale, which consists of not more than the funnel. However, Buchholzia is also a likely candidate. Species of this genus have oesophageal appendages and intestinal diverticula, and these are sufficiently similar to the respective structures in Enchylea to claim a homology. Noteworthy is the dorsal insertion of the intestinal diverticula, because lateral insertion is more common in enchytraeids (genera Henlea, Enchytronia, Guaranidrilus). However, diverticula in Buchholzia are compact bodies in VIII without lateral wings and with narrow canals. Oesophageal appendages differ in details of shape (more sac-like) and insertion (laterally and more posteriorly in IV), but other details (simple tubes with wide lumen inserting in IV) are similar. Other structures differ from Enchylea: The nephridial anteseptale is with small parts of the nephridial body, testis sacs are absent, and there are two types of coelomocytes. Further peculiarities in Enchylea, not shared by species of Enchytraeus or Buchholzia, are the short and stout vasa deferentia (present in Cernosvitoviella and some tropical Achaeta species), and the anterior location of the first nephridium (4 / 5, present also in some species of Henlea and Bryodrilus). Enchytraeus and Buchholzia occupy different branches in the molecular phylogenetic trees generated by Erséus et al (2010). DNA has been extracted from one of the three specimens of Enchylea that we found, and future sequencing may help to clarify the phylogenetic relationships of Enchylea. Enchylea heteroducta was found in low numbers (3 specimens among thousands of enchytraeids), which means that the preferred habitat type of the species may differ from the types represented by the sites sampled in this study. The high number of oocytes mentioned in the original description (up to 8 per individual) and its first appearance in laboratory cultures suggests a lifestyle similar to that of other easily culturable enchytraeid species, all of them belonging to Enchytraeus, r-strategists adapted to exploiting high amounts of all kinds of organic material. We therefore believe that higher densities of this interesting species will be found in mediterranean compost heaps.	en	Schmelz, Rüdiger M., Collado, Rut (2013): Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963. Zootaxa 3647 (2): 307-328, DOI: 10.11646/zootaxa.3647.2.4
